﻿Preliminary checklist of the genus Festuca L. (Loliinae, Pooideae, Poaceae) in the Altai Mountains with outlines for further studies

﻿Abstract Here we present an updated checklist of the genus Festuca in the Altai Mountains (AM). The study was carried out on the abundant herbarium material and considered the latest published phylogenetic analyses. Festuca was revised within the scope of the fine-leaved group (clade) with two sections, sect. Aulaxyper and sect. Festuca. Two species, namely F.richardsonii and F.lenensis, were previously misidentified and are not present in the AM. Festucabrevissima is a new record for the Russian part of the AM and for the flora of Mongolia. In total, our revision shows that 17 species of fine-leaved fescues are present in the area of AM. In this paper, we provide a key to species identification, as well as illustrations of plants, habits, leaves, spikelets, and glumes. Information on nomenclature types, synonymy, flowering period, chromosome numbers, habitats, and general distribution along with distribution maps of the particular species within the AM are included.


Introduction
The fescue genus, Festuca L., is one of the largest genera of the Poaceae family and includes more than 600 species with the greatest diversity in the Holarctic zone of Eurasia and North America (Tzvelev 1976;Alexeev 1980;Clayton and Renvoize 1986;Aiken and Darbyshire 1990;Watson and Dallwitz 1992;Soreng et al. 2003Soreng et al. , 2022;;Сlayton et al. 2006;Lu et al. 2006;Stančik and Peterson 2007).The genus is easily recognisable by its perennial caespitose or rhizomatous plant form, paniculate inflorescences, 3-11-floret spikelets, dorsally rounded lemmas with 3-5 veins, a linear hilum, and lack of a hairy appendage on the ovary apex (Kellogg 2015;Ospina-González et al. 2015).A base chromosome number of the genus Festuca is x=7 (Ospina-González et al. 2015).However, fescues are often polyploids, and even representatives of one species may be characterised by a different number of chromosomes (Šmarda and Kočí 2003;Šmarda et al. 2005;Galli et al. 2006;Enushchenko and Probatova 2020;Kiedrzyński et al. 2021).In the Altai Mountains (AM), the fescues are dip-PhytoKeys 234: 229-274 (2023), DOI: 10.3897/phytokeys.234.105385Polina D. Gudkova et al.: Checklist of the genus Festuca L. in the Altai Mountain Country loids, tetraploids or hexaploids.Different ploidy level is typical for species such as F. rubra L., F. kryloviana Reverd., F. pseudovina Hach.ex Wiesb., F. lenensis Drobow and F. valesiaca Schleich. ex Gaudin (Mizianty and Pawlus 1984;Alexeev et al. 1988;Šmarda 2008;Dirihan et al. 2016;Tzvelev and Probatova 2019).It is considered that ploidy level influences the morphological characters including the diagnostic ones, such as the length of the lemma, the length of the spikelet, the width of the vegetative leaves in the way that higher ploidy levels results in generally larger sizes of the particular parts of the plants (Litardière 1923;Levitskii and Kuzmina 1927;Bidault 1968;Rewicz et al. 2018).Therefore, the species whose representatives are characterised by polyploidy are highly polymorphic, which significantly complicates species identification.In the Altai Mountains, the fescues are diploids, tetraploids or hexaploids.Although fescues are widely distributed in the steppe zone and have regional economic importance as forage grasses, due to taxonomic problems and challenging species identifications, the genus is understudied, and thus all the more important (Torrecilla and Catalán 2002;Torrecilla et al. 2003Torrecilla et al. , 2013;;Ospina-González et al. 2015;Bednarska and Brazauskas 2017;Ospina and Picca 2019;Enushchenko and Probatova 2020;Boeuf et al. 2022;Vasile et al. 2022).
The Altai Mountains (AM) are located in Russia, Kazakhstan, Mongolia and China.The AM host 2700 species of vascular plants, with Poaceae being one of the most widely distributed families (Kamelin 2005; http://altaiflora.asu.ru/en/;Vaganov et al. 2019) including several species of fescues.The first person who listed species of Festuca in the AM was Gmelin (1747).Later, Trinius (1829) recorded six species of Festuca that belong to subgenera Festuca, Leucopoa (Griseb.)Hack., and Schedonorus Peterman.While the above mentioned authors used only macromorphological characters of flowering, and less often vegetative, shoots for species-identification, Hackel (1882) proposed using anatomical structures of the leaf blade cross-section as diagnostic characters, and these have been shown to be very significant for the taxonomy of the genus.Following Drobow (1915), botanists used these characters for the description and identification of the Festuca species in the Altai territory (Krylov 1914;Reverdatto 1936Reverdatto , 1965;;Tzvelev 1976;Grubov 1982;Alexeev 1990;Chen et al. 2003;Lu et al. 2006).
According to the results of our molecular studies based on genome-wide genotyping (Kriuchkova et al. 2023), the Altai fescues are divided into five well-supported clades.However, relationships of some species within clades (e.g.F. ovina, F. kryloviana and F. valesiaca) are weakly resolved.There are also no general findings regarding the diversity of fescue species in the AM based on a comparison of the morphology and the molecular data.Therefore, a thorough study of Festuca in the AM, including the analysis of its morphological variability corresponding to molecular data, is necessary.
The goal of this paper is to evaluate the morphological and anatomical characteristics of particular species of fine-leaved fescues in the AM.We then present an identification key, detailed distribution maps of the examined species, and illustrations of the most important characters of the examined species.

Study area
The study was carried out in 2017-2021 in the AM, which straddles the territories of Russia (South Siberia), eastern Kazakhstan, western China (Xinjiang Uygur Autonomous Region), and a part of western Mongolia, and occupies an area of 550,000 km 2 .The elevation varies from 240 m to the highest mountains of North Asia, namely Belukha (4506 m), and Tavan-Bogdo-Ula (4374 m).The AM include five elevational zones: steppe, forest-steppe, forest, subalpine, and alpine tundra.However, depending on the region, the composition and altitudinal boundaries of these zones are different.The AM host 2700 species of vascular plants, and the most widely distributed families are Asteraceae, Poa-ceae, and Fabaceae, which characterise the boreal flora (Fig. 1; Kamelin 2005; http://altaiflora.asu.ru/en/;Vaganov et al. 2019).For the AM, Kamelin (2005) proposed a botanical-geographical subdivision into regions, which we used to describe the distribution of species.

Plant material
This study is based on the revision of the specimens deposited in the following herbaria: ALTB, AA, LE, KRA, KUZ, MW, NS, NSK, and TK.In addition, we conducted a series of field work expeditions into the Russian, Mongolian and Khazakh parts of the Altai Mountains.Herbarium acronyms follow Thiers (2023, continuously updated http://sweetgum.nybg.org/ih/).All specimens were compared with the type specimens and protologues.Photos for illustration were taken with a stereo microscope Nikon SMZ800N (Japan), the image of the general habit was taken with a scanner HP Laser Jet M1132 (USA).Maps were generated using DIVA-GIS 7.5 (Hijmans et al. 2001, continuously updated http://www.diva-gis.org).In total, more than 800 herbarium sheets of 23 species were used for the preparation of distribution maps.The distribution frequency scale is as follows: very rare -1-10 localities, rare -10-20 localities, common -20-30 localities, wide spread ->50 localities.

Taxonomic treatment
Identification key to Festuca species occurring in the AM Type.Lectotype (designated by Jarvis et al. 1987: 302).Habitat in Europae sterilibus siccis (GB).General distribution.Widespread across North America, Eurasia, Africa, introduced in Australia and South America.
Distribution in the AM.Widespread, all regions (Fig. 3f).
Notes.Festuca rubra s.l. is an easily recognised taxon that has mostly extravaginal vegetative shoots (plants usually rhizomatous), the sheaths of the tillers and young leaves on flowering culms being fused almost to the top, 4-6 angular shape of leaf blades in cross-section, five or seven sclerenchyma strands, and well-defined ribs.Festuca rubra is a polymorphic species: the plant length is 25-100 cm; the colour of spikelets is greenish to purplish; the surface of lemma is characterised as glabrous or smooth with hooks, prickles or pilose, and the palea keels have hooks or prickles, the number of flowers in a spikelet is 5 to 11; the number of vascular bundles in a leaf blade cross-section of vegetative shoots equals to 5-11; the lemma length is 3.7-7 mm.Recent molecular research revealed that all of the examined F. rubra s.l.specimens belong to one clade (Kriuchkova et al. 2023).
Festuca richardsonii was previously recorded for the AM.However, these specimens were misidentified with F. rubra.Moreover, the results of our molecular study confirmed that the above mentioned specimens did not show any significant molecular differences (Kriuchkova et al. 2023), thus we here treat PhytoKeys 234: 229-274 (2023) the specimens collected from the AM and previously determined as F. richardsonii as representatives of F. rubra.
Festuca richardsonii is a species also belonging to sect.Aulaxyper.The species has a rather complicated history of records in the study area.Alexeev (1990) was the first who reported F. rubra subsp.arctica (=F.richardsonii) in the AM.He noted that, in the territory of the AM, there were individuals with transitional characters between F. rubra subsp.arctica and F. rubra subsp.rubra.A quarter of a century later, Chusovljanov (2007) recorded three more localities of F. richardsonii from the AM.Generally, F. richardsonii is the most widely distributed grass throughout the Arctic, common throughout Eurasia, North America, and Greenland.It was also recorded in the northern boreal zones, and in the Tarbagatai, Tian Shan, and Pamiro-Alai mountains within Central Asia (Markgraf-Dannenberg 1980;Darbyshire and Pavlick 2007;Chusovljanov 2007).Typical specimens of F. richardsonii differ from F. rubra in having shorter culm length (10-30(40) cm vs 50-60 cm); shape of the panicle (contracted vs open); smaller number of spikelets on lower panicle branches (1-2 spikelets vs 2 and more spikelets); the shorter awn length (0-1.5 mm vs 1-2.3 mm), respectively.
Festuca albifolia morphologically is also close to F. valesiaca.However, F. albifolia is distinguished from F. valesiaca by shoots (grouped by 2-3, surrounded by a cover of the old sheath vs single, not grouped), the leaf blade width (0.6-0.8 mm vs (0.35)0.4-0.6 mm), the number of ribs in a leaf blade cross-section (1 well-defined midrib vs 3 well-defined ribs), the number of vascular bundles (7 vs 5), the leaf sheaths of tillers (fused to ¼-¹⁄3 their length vs fused to ¹⁄6-¼ their length), the lemma length (4-5 mm vs (2.8)   Notes.Festuca borissii is close to F. kryloviana.Some taxonomists have distinguished the species from each other based on characters of the abaxial surface of the leaf blade and panicle branches, but these, in fact, cannot be used for this purpose (Tzvelev 1976;Alexeev 1990;Tzvelev and Probatova 2019).Both species are characterised by glabrous or scabrous leaf blades, scabrous or pubescent panicle branches.Festuca borissii differs from F. kryloviana by the structure of tillers (loosely tufted with extravaginal shoots vs densely tufted with intravaginal shoots respectively), the shape of the leaf blade cross-section (elongated  Chromosome number.2n=42 (Kamchatskaya obl., Magadanskii obl.; Alexeev et al. 1987b;o. Vrangelya;Zhukova and Petrovskii 1980;Chukotka;Yurtzev and Zhukova 1978;American Arctica;Frederiksen 1981) Notes.Festuca brachyphylla is easily recognised by its panicles with more than 2 spikelets on lower branches, anthers 0.5-1.5 mm in length, scabrid leaf blades, 5 or 7 sclerenchyma strands.Phylogenetically, our molecular study demonstrated that F. brachyphylla and F. brevissima refer to a separate clade distinct from sect.Festuca (Kriuchkova et al. 2023 Notes.Festuca brevissima is distributed in Chukotka, Kamchatka, Wrangel Island and Alaska on gravel slopes.As a result of the herbarium revision, we recorded the species much farther to the South in the mountains of Siberia.The specimens have been misidentified as F. brachyphylla.Festuca brevissima differs from F. brachyphylla by the number of spikelets in the panicle (fewer than 8 spikelets vs more than 11 spikelets respectively), the number of spikelets on lower branches (1-2 spikelets vs 2-or more spikelets), the panicle length (0.7-26 mm vs 23-55 mm), the lemma length (2.5-4 mm vs 4.5-5.5 mm), the plant length (up to 120 mm vs 100-550 mm).According to the results of our molecular studies, F. brevissima is clearly separated from F. brachyphylla (Kriuchkova et al. 2023).All the localities listed above are new records of the species for the AM.Information on the occurrence of species in areas previously unnoticed or misidentified with other species is nowadays very important, because being under increasing pressure from human activities and recently also under  General distribution.China (Altai, Dzungaria), Mongolia (Altai) and Russia (Altai, Sayan, Ural, Tarbagatai), Kazakhstan (Altai, Tyan-Shan,).
Notes.Festuca kryloviana is a morphologically variable species.The surface of lemma can be glabrous and smooth or scabrous, abaxial surface of the vegetative leaf blade is glabrous to scabrous, adaxial surface of the vegetative leaf blade is covered by hooks or prickles, rarely with microhairs present, the number of vascular bundles is 5 to 7, rarely 9 and the number of ribs is 3 to rarely 5, the lemma length equals to 4.3-5 mm, the awn length varies from 2 to 5 mm; the wax on the leaf blades is present or absent.
Notes.The distribution of Festuca kurtschumica is restricted to the AM.Festuca kurtschumica is similar to F. kryloviana but differs by leaf sheaths of tillers (fused for ²⁄3-¾ their length vs fused for ¹⁄3-½ their length), leaf blade width (0.4-0.5 (0.55) mm vs (0.4)0.55-0.85(1)mm), lemma length (3.5-4.5 mm vs 4.5-6 mm), and the number of vascular bundles (5 vs (5)7 respectively).However, in accordance with the results of molecular analysis, F. kurtschumica and F. kryloviana are grouped into a common clade (Kriuchkova et al. 2023).The species needs a taxonomic revision within its entire distributional range, including the locus classicus, namely the vicinity of Lake Markakol, the Khazakh part of the AM.
Specimens examined.Russia.General distribution.The species occurs in mountains, from Turkey, throughout Caucasus, Central Asia (Western, Eastern Siberia), up to Mongolia (Altai).

Festuca ovina agg.
The aggregate comprises three species in the AM, F. ovina, F. sphagnicola and F. kuprijanovii.
Notes.Festuca ovina is easily distinguished by green or bluish green spikelets, the leaf blade cross-section with abaxial sclerenchyma in a continuous or sometimes discontinuous in 3 main islets of low profile layer, with only a midrib well defined, and with 5 or 7 vascular bundles.Within the F. ovina complex in the territory of the AM, two more species close to F. ovina (F.sphagnicola and F. kuprijanovii) are identified, however molecularly they form a common clade (Kriuchkova et al. 2023).
The similar issue refers to another species of the F. ovina complex, F. sphagnicola.Some botanists treated F. sphagnicola as a subspecies of F. ovina (Tzvelev 1971(Tzvelev , 1976;;Probatova and Sokolovskaya 1980;Alexeev 1990;Lu et al. 2006), or independent species (Tzvelev and Probatova 2019).Further, Alexeev considered diploid individuals to be F. ovina s. str.and referred polyploid individuals to other species.Consequently, tetraploid individuals of F. ovina from the AM were referred to F. sphagnicola (Table 1).The last mentioned taxon differs from F. ovina by the color of spikelets (brown vs green, respectively), the elevation of occurrence (over 1700 m vs up to 1700 m, respectively), the number of chromosomes (tetraploid vs diploid).Molecular research, however, grouped F. sphagnicola, F. kuprijanovii and F. ovina into a common clade (Kriuchkova et al. 2023).Thus, further studies including molecular (at the population level), morphological, and cytological analyses are needed on the above mentioned group of taxa from the entire area of their distribution to resolve whether F. kuprijanovii and F. sphagnicola are separate or conspecific with F. ovina.General distribution.The species is widely distributed, it is common in the Arctic zone of Eurasia and North America, and occurs also in mountains farther to the south of Eurasia (Kazakhstan, China, Mongolia).
Notes.Festuca pseudovina is morphologically similar and closely related to F. valesiaca.The color of the leaves (bluish-green or green) is the only morphological character that separates these species (Alexeev 1990;Tzvelev 1976;Lu et al. 2006;Tzvelev and Probatova 2019).Most specimens previously identified as F. pseudovina have been re-identified by us as F. valesiaca, because both are bluish green but are characterised by varying colour saturation of leaf blades.In molecular research, F. pseudovina specimens collected from the AM were shown to be hybrids between F. valesiaca and F. rupicola (Kriuchkova et al. 2023).The species needs further study, including molecular (at the population level), morphological, and cytological analyses.
General distribution.Saur ridge, endemic of Altai.Distribution in the AM.Very rare; KAD4 (Fig. 15d).Habitat.Forests, alpine zone; elev.2000-2600 m.Flowering period.June-July.Chromosome number.2n=unknown.Notes.Festuca saurica is an endemic species of the AM.The species is highly variable in morphology: the number of vascular bundles varies from 5 to 7; the leaf blade cross-section may have only one single midrib or two additional lateral well-defined ribs; the shape of ribs varies from rounded to triangular; the shape of the leaf blade cross-section varies from obovate with an elongated keel to wide-lanceolate; the leaf sheaths of tillers may be fused for ½-¾ their length; the abaxial surface of the leaf blade is characterised as glabrous or scabrous; and shoots are either grouped by 2-3 and surrounded by a cover of old sheaths.According to our molecular research, Festuca saurica is separated in an independent clade (Kriuchkova et al. 2023).
Notes.The species is easily distinguished by its lemmas 4.5-6 mm long, awns 1.5-4 mm long, leaf blades 0.5-1.1 mm wide, leaf blade cross-section with 3 well-defined ribs, leaf sheaths of tillers fused for ¹⁄3-½ their length; shoots are either grouped by 2-3 and surrounded by a cover of old sheaths.Our recent molecular research revealed that this species might hybridise with the F. kryloviana group (Kriuchkova et al. 2023).However, there are no morphological characters distinguishing the hybrids from F. tschujensis.Alexeev (1981) and Chusovljanov (2007) mention that F. lenensis grows in the Mongolian part of the AM, and the species differs from F. tschujensis by the leaf blade structure (width of middle sclerenchyma strand is similar to lateral strands vs middle sclerenchyma strand is two-three times wider than lateral strands; leaf blades arcuate vs flexuose, respectively).However, during our revision, all the available specimens of F. lenensis from the AM were identified by the first author of this paper as F. tschujensis.Thus, the presence of F. lenensis in the AM needs to be confirmed.F. valesiaca includes two genetic groups: one from mountains and another from lowland (Kriuchkova et al. 2023).The species needs further study, including molecular (at the population level), morphological, and cytological analyses.

Conclusion
During our study we confirmed the occurrence of 17 species of genus Festuca in the AM.Festuca brevissima is a new record to the flora of the AM and at present, its localities in the region are the southernmost within its distribution range.Our revision showed that F. lenensis and F. richardsonii are not component of the AM flora.All the available specimens of F. lenensis from the AM were identified as F. tschujensis, whereas specimens of F. richardsonii previously recorded for the AM were identified as F. rubra.We found also that F. oreophylla was previously listed in the flora of the AM as a synonym of F. musbelica, however, F. musbelica is the earlier legitimate name.Finally, we did not confirm Chusovljanov's assumption (Chusovljanov 2003) on the presence of F. kemerovensis in the AM.
In accordance with our morphological and molecular (Kriuchkova et al. 2023) studies we have revealed three complexes of closely related taxa: F. kurtschumica-F. kryloviana; F. sphagnicola-F. kuprijanovii-F. ovina; and F. musbelica-F. valesiaca.Nevertheless, to resolve whether they are separate or conspecific species, further studies including more advanced molecular, morphological, and cytological analyses are needed on the above mentioned complexes of taxa from the entire area of their distribution.

Figure 2 .
Figure 2. The most important macromorphological and anatomical characters of fescues A, B, I a cross-section with three well-defined ribs C, D, G, H a cross-section with only midrib or also two lateral ribs weakly defined.Shape of the cross-section leaf blades: A 4 angular B, C, H obovate A, C, D diagram of the leaf blades' anatomical structure of Festuca (designed by E.B. Alexeev): а) diameter, b/c) ratio b/c, 1) lateral sclerenchyma strand, 2) lateral ribs, 3) keel sclerenchyma strand, 4) middle rib, 5) vascular bundles, 6) continuous sclerenchyma layer C lateral sclerenchyma strands similar middle strand H lateral sclerenchyma strands less, than middle strand F flower: a) awn, b) lemma E panicle with 1-2 spikelets on lower branches, contacted J panicle with more than 2 spikelets on lower branches, open K grouped shoots (2-3 shoots surrounded by old sheaths) L single shoots: 1) flowering shoot, 2) vegetative shoot (AKA tillers).

Figure 3 .
Figure 3. Festuca rubra a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species distribution in the region known from literature data; red points mark localities confirmed by us during the revision of herbarium materials.

Figure 4 .
Figure 4. Festuca albifolia a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map marks information about distribution for the region known from literature data, red points mark localities confirmed by herbarium materials revised during our studies.

Figure 5 .
Figure 5. Festuca borissii a spikelet, lateral view b glumes, lateral view c leaf-blade cross-section d junction of leaf sheath and blade, lateral view e distribution map f general habit.Scale bars: 10 cm (f); 1 mm (a, b, d); 0.5 mm (c).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 6 .
Figure 6.Festuca brachyphylla a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, while red points mark localities confirmed by us during revision of herbarium materials.

Figure 7 .
Figure 7. Festuca brevissima a general habit b junction of leaf sheath and blade, lateral view c spikelet, lateral view d glumes, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).

Figure 8 .
Figure 8. Festuca kryloviana a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 9 .
Figure 9. Festuca kurtschumica a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 10 .
Figure 10.Festuca musbelica a general habit b spikelet, lateral view c distribution map d glumes, lateral view e leaf-blade cross-section f junction of leaf sheath and blade, lateral view.Scale bars: 10 cm (a); 1 mm (b, d, f); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 11 .
Figure 11.General habit of a F. ovina b F. kuprijanovii j F. sphagnicola glumes, lateral view c F. sphagnicola d F. ovina spikelets e F. sphagnicola f F. ovina junction of leaf sheath and blade, lateral view g F. sphagnicola h F. ovina i F. kuprijanovii leaf-blade cross-section l F. sphagnicola (two cross-sections) m F. ovina n F. kuprijanovii distribution maps k F. sphagnicola o F. ovina p F. kuprijanovii.Scale bars: 10 cm (a, b, j); 1 mm (c-i); 0.5 mm (l-n).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 12 .
Figure 12.Festuca pseudovina a general habit b glumes, lateral view c spikelet, lateral view d junction of leaf sheath and blade, lateral view e leaf-blade cross-section f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 13 .
Figure 13.Festuca pseudosulcata a general habit b glumes, lateral view c leaf-blade cross-section d junction of leaf sheath and blade, lateral view e spikelet, lateral view f distribution map.Scale bars: 10 cm (a); 1 mm (b, d, e); 0.5 mm (c).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 15 .
Figure 15.Festuca saurica a general habit b distribution map of the AM c glumes, lateral view d spikelet, lateral view e junction of leaf sheath and blade, lateral view f leaf-blade cross-section.Scale bars: 10 cm (a); 1 mm (c-e); 0.5 mm (f).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 16 .
Figure 16.Festuca tschujensis a general habit b distribution map of the AM c glumes, lateral view d spikelet, lateral view e junction of leaf sheath and blade, lateral view f leaf-blade cross-section.Scale bars: 10 cm (a); 1 mm (c-e); 0.5 mm (f).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Figure 17 .
Figure 17.Festuca valesiaca a general habit b spikelet, lateral view c glumes, lateral view d leaf-blade cross-section e junction of leaf sheath and blade, lateral view f distribution map.Scale bars: 10 cm (a); 1 mm (b-d); 0.5 mm (e).The green colour on the map refers to information on species-distribution in the region known from literature data, red points mark localities confirmed by us during revision of herbarium materials.

Table 1 .
Main morphological differences between species of the F. ovina aggregate.
Notes.See notes under Festuca agg ovina.